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Quaternary Vertebrate Faunas from Sumba, Indonesia: Implications for Wallacean Biogeography and Evolution Samuel Turvey University of Cambridge

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Quaternary Vertebrate Faunas from Sumba, Indonesia: Implications for Wallacean Biogeography and Evolution Samuel Turvey University of Cambridge University of Wollongong Research Online Faculty of Science, Medicine and Health - Papers Faculty of Science, Medicine and Health 2017 Quaternary vertebrate faunas from Sumba, Indonesia: Implications for Wallacean biogeography and evolution Samuel Turvey University of Cambridge Jennifer Crees University of Cambridge James Hansford University of Southampton Timothy Jeffree University of Cambridge Nick Crumpton University of Cambridge See next page for additional authors Publication Details Turvey, S. T., Crees, J. J., Hansford, J., Jeffree, T. E., Crumpton, N., Kurniawan, I., Setiyabudi, E., Guillerme, T., Paranggarimu, U., Dosseto, A. & Van Den Bergh, G. D. (2017). Quaternary vertebrate faunas from Sumba, Indonesia: Implications for Wallacean biogeography and evolution. Proceedings of the Royal Society B: Biological Sciences, 284 (1861), 20171278-1-20171278-10. Research Online is the open access institutional repository for the University of Wollongong. For further information contact the UOW Library: [email protected] Quaternary vertebrate faunas from Sumba, Indonesia: Implications for Wallacean biogeography and evolution Abstract Historical patterns of diversity, biogeography and faunal turnover remain poorly understood for Wallacea, the biologically and geologically complex island region between the Asian and Australian continental shelves. A distinctive Quaternary vertebrate fauna containing the small-bodied hominin Homo floresiensis, pygmy Stegodon proboscideans, varanids and giant murids has been described from Flores, but Quaternary faunas are poorly known from most other Lesser Sunda Islands. We report the discovery of extensive new fossil vertebrate collections from Pleistocene and Holocene deposits on Sumba, a large Wallacean island situated less than 50 km south of Flores. A fossil assemblage recovered from a Pleistocene deposit at Lewapaku in the interior highlands of Sumba, which may be close to 1 million years old, contains a series of skeletal elements of a very small Stegodon referable to S. sumbaensis, a tooth attributable to Varanus komodoensis, and fragmentary remains of unidentified giant murids. Holocene cave deposits at Mahaniwa dated to approximately 2000-3500 BP yielded extensive material of two new genera of endemic large-bodied murids, as well as fossils of an extinct frugivorous varanid. This new baseline for reconstructing Wallacean faunal histories reveals that Sumba's Quaternary vertebrate fauna, although phylogenetically distinctive, was comparable in diversity and composition to the Quaternary fauna of Flores, suggesting that similar assemblages may have characterized Quaternary terrestrial ecosystems on many or all of the larger Lesser Sunda Islands. Disciplines Medicine and Health Sciences | Social and Behavioral Sciences Publication Details Turvey, S. T., Crees, J. J., Hansford, J., Jeffree, T. E., Crumpton, N., Kurniawan, I., Setiyabudi, E., Guillerme, T., Paranggarimu, U., Dosseto, A. & Van Den Bergh, G. D. (2017). Quaternary vertebrate faunas from Sumba, Indonesia: Implications for Wallacean biogeography and evolution. Proceedings of the Royal Society B: Biological Sciences, 284 (1861), 20171278-1-20171278-10. Authors Samuel Turvey, Jennifer Crees, James Hansford, Timothy Jeffree, Nick Crumpton, Iwan Kurniawan, Erick Setiyabudi, Thomas Guillerme, Umbu Paranggarimu, Anthony Dosseto, and Gerrit D. van den Bergh This journal article is available at Research Online: http://ro.uow.edu.au/smhpapers/4954 Downloaded from http://rspb.royalsocietypublishing.org/ on September 18, 2017 Quaternary vertebrate faunas from rspb.royalsocietypublishing.org Sumba, Indonesia: implications for Wallacean biogeography and evolution Samuel T. Turvey1, Jennifer J. Crees1, James Hansford1,2, Timothy E. Jeffree1, Research Nick Crumpton1, Iwan Kurniawan3, Erick Setiyabudi3, Thomas Guillerme4, Cite this article: Turvey ST et al. 2017 Umbu Paranggarimu5, Anthony Dosseto6 and Gerrit D. van den Bergh7 Quaternary vertebrate faunas from Sumba, 1Institute of Zoology, Zoological Society of London, Regent’s Park, London NW1 4RY, UK Indonesia: implications for Wallacean 2Ocean and Earth Science, National Oceanography Centre, University of Southampton, Southampton SO14 3ZH, UK biogeography and evolution. Proc. R. Soc. B 3Geology Museum Bandung, Bandung 40122, Indonesia 284: 20171278. 4Department of Life Sciences, Imperial College London, Silwood Park, Ascot SL5 7PY, UK 5 http://dx.doi.org/10.1098/rspb.2017.1278 KOPPESDA, Kampung Arab, Waingapu, Sumba Timur, Indonesia 6Wollongong Isotope Geochronology Laboratory, School of Earth and Environmental Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia 7Centre for Archaeological Science, School of Earth and Environmental Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia Received: 7 June 2017 STT, 0000-0002-3717-4800; TG, 0000-0003-4325-1275 Accepted: 21 July 2017 Historical patterns of diversity, biogeography and faunal turnover remain poorly understood for Wallacea, the biologically and geologically complex island region between the Asian and Australian continental shelves. A distinc- Subject Category: tive Quaternary vertebrate fauna containing the small-bodied hominin Homo Palaeobiology floresiensis, pygmy Stegodon proboscideans, varanids and giant murids has been described from Flores, but Quaternary faunas are poorly known from most other Lesser Sunda Islands. We report the discovery of extensive new Subject Areas: fossil vertebrate collections from Pleistocene and Holocene deposits on palaeontology Sumba, a large Wallacean island situated less than 50 km south of Flores. A fossil assemblage recovered from a Pleistocene deposit at Lewapaku in the Keywords: interior highlands of Sumba, which may be close to 1 million years old, contains biogeography, island evolution, murid, a series of skeletal elements of a very small Stegodon referable to S. sumbaensis, Quaternary extinctions, Stegodon, Wallacea a tooth attributable to Varanus komodoensis, and fragmentary remains of un- identified giant murids. Holocene cave deposits at Mahaniwa dated to approximately 2000–3500 BP yielded extensive material of two new genera of endemic large-bodied murids, as well as fossils of an extinct frugivorous vara- Authors for correspondence: nid. This new baseline for reconstructing Wallacean faunal histories reveals Samuel T. Turvey that Sumba’s Quaternary vertebrate fauna, although phylogenetically distinc- e-mail: [email protected] tive, was comparable in diversity and composition to the Quaternary fauna of Gerrit D. van den Bergh Flores, suggesting that similar assemblages may have characterized Quaternary terrestrial ecosystems on many or all of the larger Lesser Sunda Islands. e-mail: [email protected] 1. Introduction Islands represent important ‘natural laboratories’ for investigating evolutionary patterns, processes and dynamics [1,2], and unusual patterns of morphological evolution are observed in island environments, including gigantism in small- bodied lineages and dwarfism in large-bodied lineages [3–5]. However, insular taxa are particularly vulnerable to human pressures, and many island systems have lost a substantial proportion of their Quaternary vertebrate faunas following prehistoric human arrival [6,7]. Reconstructing historical diversity is therefore essential to overcome this ‘extinction filter’ and understand the evolutionary origin, morphological adaptation and biogeographic affinity of island faunas. Electronic supplementary material is available Such research is particularly important in southeast Asia, a geologically com- online at https://dx.doi.org/10.6084/m9. plex region with spatially complex biodiversity and endemism, which has figshare.c.3844780. & 2017 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited. Downloaded from http://rspb.royalsocietypublishing.org/ on September 18, 2017 2 119° E 120° E rspb.royalsocietypublishing.org 2 1 Proc. R. Soc. B 10° S 3 284 Bali : 20171278 Flores Lombok Sumbawa Timor Sumba N 50 km 50 km Figure 1. Quaternary fossil localities on Sumba. 1, Lewapaku; 2, Watumbaka; 3, Mahaniwa. Inset, location of Sumba in Lesser Sundas, showing direction of Indo- nesian Throughflow past Flores and nearby islands towards Sumba. provided insights into biogeographic processes for over a cen- Flores has never been joined to other large islands in the tury [1]. In particular, faunal relationships across Wallacea, Lesser Sundas, but its extant fauna shows biogeographic affi- the islands that were never connected with the Asian or Austra- nities with nearby islands [8,9], suggesting that ecologically lian continental shelves during periods of low sea level, similar Quaternary vertebrate faunas may also have had a challenge easy interpretation [8–10]. Some vertebrate groups wider former distribution across Wallacea. Distinctive extinct (e.g. proboscideans) are known to have become completely Quaternary vertebrates (mainly endemic Stegodon species, extinct in Wallacea during the Late Quaternary, possibly rodents and/or other small-bodied vertebrate taxa) have also owing to prehistoric interactions with human colonists [4,5] or been described from Sulawesi, Timor, Halmahera and some alternatively because of climatic factors or postulated long- smaller Wallacean islands [4,10,13,29–34]. In particular, two term
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